Please note that I have moved to the University of South Carolina, as Director of the School of the Earth, Ocean & Environment and Professor of Biological Sciences.

Academic Appointments

Administrative Appointments

  • Bing Director, Program in Human Biology (2006 - 2012)

Honors & Awards

  • Fellow, California Academy of Sciences (2000)
  • Fellow, American Association for the Advancement of Science (2001)
  • Stanford Friends University Fellow in Undergraduate Education, Stanford University (2010-)

Professional Education

  • PhD, University of Texas at Austin, Zoology (1979)

Current Research and Scholarly Interests


We are exploring how environmental variation affects life history traits, population structure and dynamics, and species interactions in ecological and evolutionary time, using Lepidoptera. Current interests include (1) how resource allocation strategies interact with foraging and life history in variable environments to affect fitness and population dynamics; (2) the ecological and evolutionary dynamics of small populations, including population re-introductions; and (3) invasion biology, particularly the evolutionary and ecological effects of non-native species' invasion into co-evolved systems.

2017-18 Courses

Graduate and Fellowship Programs

  • Biology (School of Humanities and Sciences) (Phd Program)

All Publications

  • The fingerprints of global climate change on insect populations CURRENT OPINION IN INSECT SCIENCE Boggs, C. L. 2016; 17: 69-73


    Synthesizing papers from the last two years, I examined generalizations about the fingerprints of climate change on insects' population dynamics and phenology. Recent work shows that populations can differ in response to changes in climate means and variances. The part of the thermal niche occupied by an insect population, voltinism, plasticity and adaptation to weather perturbations, and interactions with other species can all exacerbate or mitigate responses to climate change. Likewise, land use change or agricultural practices can affect responses to climate change. Nonetheless, our knowledge of effects of climate change is still biased by organism and geographic region, and to some extent by scale of climate parameter.

    View details for DOI 10.1016/j.cois.2016.07.004

    View details for Web of Science ID 000386420900012

    View details for PubMedID 27720076

  • Nutrient acquisition across a dietary shift: fruit feeding butterflies crave amino acids, nectivores seek salt OECOLOGIA Ravenscraft, A., Boggs, C. L. 2016; 181 (1): 1-12


    Evolutionary dietary shifts have major ecological consequences. One likely consequence is a change in nutrient limitation-some nutrients become more abundant in the diet, others become more scarce. Individuals' behavior should change accordingly to match this new limitation regime: they should seek out nutrients that are deficient in the new diet. We investigated the relationship between diet and responses to nutrients using adult Costa Rican butterflies with contrasting feeding habits, testing the hypothesis that animals will respond more positively to nutrients that are scarcer in their diets. Via literature searches and our own data, we showed that nitrogen and sodium are both at lower concentration in nectar than in fruit. We therefore assessed butterflies' acceptance of sodium and four nitrogenous compounds that ranged in complexity from inorganic nitrogen (ammonium chloride) to protein (albumin). We captured wild butterflies, offered them aqueous solutions of each substance, and recorded whether they accepted (drank) or rejected each substance. Support for our hypothesis was mixed. Across the sexes, frugivores were four times more likely to accept amino acids (hydrolyzed casein) than nectivores, in opposition to expectation. In males, nectivores accepted sodium almost three times more frequently than frugivores, supporting expectations. Together, these results suggest that in butterflies, becoming frugivorous is associated with an increased receptivity to amino acids and decreased receptivity to sodium. Nectivory and frugivory are widespread feeding strategies in organisms as diverse as insects, birds, and bats; our results suggest that these feeding strategies may put different pressures on how animals fulfill their nutritional requirements.

    View details for DOI 10.1007/s00442-015-3403-6

    View details for Web of Science ID 000374564000001

    View details for PubMedID 26267402

  • Effects of larval dietary restriction on adult morphology, with implications for flight and life history ENTOMOLOGIA EXPERIMENTALIS ET APPLICATA Boggs, C. L., Niitepold, K. 2016; 159 (2): 189-196

    View details for DOI 10.1111/eea.12420

    View details for Web of Science ID 000375076200009

  • Fine-Grained Distribution of a Non-Native Resource Can Alter the Population Dynamics of a Native Consumer PLOS ONE Nakajima, M., Boggs, C. L. 2015; 10 (11)


    New interactions with non-native species can alter selection pressures on native species. Here, we examined the effect of the spatial distribution of a non-native species, a factor that determines ecological and evolutionary outcomes but that is poorly understood, particularly on a fine scale. Specifically, we explored a native butterfly population and a non-native plant on which the butterfly oviposits despite the plant's toxicity to larvae. We developed an individual-based model to describe movement and oviposition behaviors of each butterfly, which were determined by plant distribution and the butterfly's host preference genotype. We estimated the parameter values of the model from rich field data. We simulated various patterns of plant distributions and compared the rates of butterfly population growth and changes in the allele frequency of oviposition preference. Neither the number nor mean area of patches of non-native species affected the butterfly population, whereas plant abundance, patch shape, and distance to the nearest native and non-native patches altered both the population dynamics and genetics. Furthermore, we found a dramatic decrease in population growth rates when we reduced the distance to the nearest native patch from 147 m to 136 m. Thus changes in the non-native resource distribution that are critical to the fate of the native herbivore could only be detected at a fine-grained scale that matched the scale of a female butterfly's movement. In addition, we found that the native butterfly population was unlikely to be rescued by the exclusion of the allele for acceptance of the non-native plant as a host. This study thus highlights the importance of including both ecological and evolutionary dynamics in analyses of the outcome of species interactions and provides insights into habitat management for non-native species.

    View details for DOI 10.1371/journal.pone.0143052

    View details for Web of Science ID 000365153400035

    View details for PubMedID 26575843

    View details for PubMedCentralID PMC4648569

  • Effects of Increased Flight on the Energetics and Life History of the Butterfly Speyeria mormonia PLOS ONE Niitepold, K., Boggs, C. L. 2015; 10 (10)

    View details for DOI 10.1371/journal.pone.0140104

    View details for Web of Science ID 000363918100019

    View details for PubMedID 26510164

  • Resource allocation as a driver of senescence: Life history tradeoffs produce age patterns of mortality JOURNAL OF THEORETICAL BIOLOGY Davison, R., Boggs, C. L., Baudisch, A. 2014; 360: 251-262
  • Resource allocation as a driver of senescence: life history tradeoffs produce age patterns of mortality. Journal of theoretical biology Davison, R., Boggs, C. L., Baudisch, A. 2014; 360: 251-262


    We investigate the effects of optimal time and resource allocation on age patterns of fertility and mortality for a model organism with (1) fixed maximum lifespan, (2) distinct juvenile and adult diets, and (3) reliance on nonrenewable resources for reproduction. We ask when it is optimal to tolerate starvation vs. conserve resources and then examine the effects of these decisions on adult mortality rates. We find that (1) age-related changes in tradeoffs partition the life cycle into as many as four discrete phases with different optimal behavior and mortality patterns, and (2) given a cost of reproduction, terminal investment can produce a signal of actuarial senescence. Also, given limitations imposed by non-replenishable resources, individuals beginning adult life with more replenishable resources do not necessarily live longer, since they can engage in capital breeding and need not defer reproduction to forage; low reproductive overheads and low costs of starvation also encourage capital breeding and may lead to earlier terminal investment and earlier senescence. We conclude that, even for species with qualitatively similar life histories, differences in physiological, behavioral and environmental tradeoffs or constraints may strongly influence optimal allocation schedules and produce variation in mortality patterns and life expectancy.

    View details for DOI 10.1016/j.jtbi.2014.07.015

    View details for PubMedID 25051533

  • Insights from Stable Isotopic Tracers on Reproductive Allocation under Stress INTEGRATIVE AND COMPARATIVE BIOLOGY Boggs, C. L., Niitepold, K. 2014; 54 (5): 880-889


    Fecundity is affected by changes in the nutritional and energetic environment, as a result of changes in acquisition, assimilation, or allocation of macro-nutrients and micro-nutrients. Stable isotopes of carbon and nitrogen offer a window into the processes underlying these changes. In insects that feed on nectar as adults, carbon isotopes can be used to trace allocation of carbon to eggs from larval (capital) and adult (income) sources. If adults are fed sugar-water, there is no source of nitrogen from the adult diet. Thus, nitrogen isotopes in eggs reflect fractionation of larval nitrogen due to protein catabolism and anabolism. We subjected adult females of two butterfly species, Speyeria mormonia and Colias eurytheme, to dietary restriction (DR), larval female S. mormonia to DR, and adult female S. mormonia to extra flight. Females subjected to extra flight were previously found to eat more as adults and to have a higher resting metabolic rate. As predicted, significantly less carbon obtained by feeding as adults was incorporated into eggs in both species under DR when adult. Speyeria mormonia eggs contained significantly more carbon derived from adult feeding under DR as larvae and when subjected to extra flight as adult females. Again as predicted, eggs from females of both species subjected to DR when adults were enriched for (15)N, suggesting that increased protein catabolism or anabolism generated additional carbon compounds. Speyeria mormonia eggs from females subjected to DR when larvae or to additional flight as adults were depleted for (15)N. The result for DR of larvae suggests minimization of protein catabolism when protein reserves are relatively scarce. The results for flight were not as predicted, and deserve further exploration. In most cases, isotopic signature in eggs changed with females' age. Eggs were progressively more enriched for the carbon signature of adults, consistent with a two-compartment mixing model for the carbon sources of larvae and adults. Eggs laid across the life of a female were progressively depleted for (15)N, followed by stabilization. This could be due to high total investment in eggs early in life, as the results are consistent with those for other growing animals. Overall, these results indicate shifts in allocation of incoming and stored (capital) carbon in response to various environmental stresses. The results for nitrogen suggest hypotheses to be tested concerning nitrogen metabolism under environmental stress.

    View details for DOI 10.1093/icb/icu074

    View details for Web of Science ID 000344601400013

    View details for PubMedID 24920750

  • From global change to a butterfly flapping: biophysics and behaviour affect tropical climate change impacts. Proceedings. Biological sciences / The Royal Society Bonebrake, T. C., Boggs, C. L., Stamberger, J. A., Deutsch, C. A., Ehrlich, P. R. 2014; 281 (1793)


    Difficulty in characterizing the relationship between climatic variability and climate change vulnerability arises when we consider the multiple scales at which this variation occurs, be it temporal (from minute to annual) or spatial (from centimetres to kilometres). We studied populations of a single widely distributed butterfly species, Chlosyne lacinia, to examine the physiological, morphological, thermoregulatory and biophysical underpinnings of adaptation to tropical and temperate climates. Microclimatic and morphological data along with a biophysical model documented the importance of solar radiation in predicting butterfly body temperature. We also integrated the biophysics with a physiologically based insect fitness model to quantify the influence of solar radiation, morphology and behaviour on warming impact projections. While warming is projected to have some detrimental impacts on tropical ectotherms, fitness impacts in this study are not as negative as models that assume body and air temperature equivalence would suggest. We additionally show that behavioural thermoregulation can diminish direct warming impacts, though indirect thermoregulatory consequences could further complicate predictions. With these results, at multiple spatial and temporal scales, we show the importance of biophysics and behaviour for studying biodiversity consequences of global climate change, and stress that tropical climate change impacts are likely to be context-dependent.

    View details for DOI 10.1098/rspb.2014.1264

    View details for PubMedID 25165769

  • Aging, Life Span, and Energetics under Adult Dietary Restriction in Lepidoptera PHYSIOLOGICAL AND BIOCHEMICAL ZOOLOGY Niitepold, K., Perez, A., Boggs, C. L. 2014; 87 (5): 684-694


    Stressful conditions can affect resource allocation among different life-history traits. The effect of dietary restriction (DR) on longevity and reproduction has been studied in many species, but we know little about its effects on energetics, especially in flying animals that have high energy demand. We assessed the effects of DR on metabolic rate throughout the entire adult life span in two butterfly species, Colias eurytheme and Speyeria mormonia. We cut the food intake of adult females in half and measured resting metabolic rate (RMR) and flight metabolic rate (FMR) together with body mass repeatedly throughout life. In both species, DR reduced body mass, but mass-corrected FMR was not affected, indicating that flight capacity was retained. DR lowered RMR and reduced fecundity but had no effect on life span. FMR declined with age, but the rate of senescence was not affected by DR. In contrast, aging had a strong negative effect on RMR only in control females, whereas food-restricted females had more stable RMR throughout their lives. The results suggest that flight capacity is conserved during nutritional stress but that investment in flight and survival may negatively affect other important physiological processes when resources are limited.

    View details for DOI 10.1086/677570

    View details for Web of Science ID 000342670000010

    View details for PubMedID 25244380

  • Genomic inference accurately predicts the timing and severity of a recent bottleneck in a nonmodel insect population. Molecular ecology McCoy, R. C., Garud, N. R., Kelley, J. L., Boggs, C. L., Petrov, D. A. 2014; 23 (1): 136-150


    The analysis of molecular data from natural populations has allowed researchers to answer diverse ecological questions that were previously intractable. In particular, ecologists are often interested in the demographic history of populations, information that is rarely available from historical records. Methods have been developed to infer demographic parameters from genomic data, but it is not well understood how inferred parameters compare to true population history or depend on aspects of experimental design. Here, we present and evaluate a method of SNP discovery using RNA sequencing and demographic inference using the program δaδi, which uses a diffusion approximation to the allele frequency spectrum to fit demographic models. We test these methods in a population of the checkerspot butterfly Euphydryas gillettii. This population was intentionally introduced to Gothic, Colorado in 1977 and has as experienced extreme fluctuations including bottlenecks of fewer than 25 adults, as documented by nearly annual field surveys. Using RNA sequencing of eight individuals from Colorado and eight individuals from a native population in Wyoming, we generate the first genomic resources for this system. While demographic inference is commonly used to examine ancient demography, our study demonstrates that our inexpensive, all-in-one approach to marker discovery and genotyping provides sufficient data to accurately infer the timing of a recent bottleneck. This demographic scenario is relevant for many species of conservation concern, few of which have sequenced genomes. Our results are remarkably insensitive to sample size or number of genomic markers, which has important implications for applying this method to other nonmodel systems.

    View details for DOI 10.1111/mec.12591

    View details for PubMedID 24237665

  • Fitness costs of butterfly oviposition on a lethal non-native plant in a mixed native and non-native plant community OECOLOGIA Nakajima, M., Boggs, C. L., Bailey, S., Reithel, J., Paape, T. 2013; 172 (3): 823-832


    Non-native plants may be unpalatable or toxic, but have oviposition cues similar to native plants used by insects. The herbivore will then oviposit on the plant, but the offspring will be unable to develop. While such instances have been described previously, the fitness costs at the population level in the wild due to the presence of the lethal host have not been quantified, for this or other related systems. We quantified the fitness cost in the field for the native butterfly Pieris macdunnoughii in the presence of the non-native crucifer Thlaspi arvense, based on the spatial distributions of host plants, female butterflies and eggs in the habitat and the survival of the larvae in the wild. We found that 2.9 % of eggs were laid on T. arvense on average, with a survival probability of 0, yielding a calculated fitness cost of 3.0 % (95 % confidence interval 1.7-3.6 %) due to the presence of the non-native in the plant community. Survival probability to the pre-pupal stage for eggs laid on two native crucifers averaged 1.6 % over 2 years. The magnitude of the fitness cost will vary temporally and spatially as a function of the relative abundance of the non-native plant. We propose that the fine-scale spatial structure of the plant community relative to the butterflies' dispersal ability, combined with the females' broad habitat use, contributes to the fitness costs associated with the non-native plant and the resulting evolutionary trap.

    View details for DOI 10.1007/s00442-012-2537-z

    View details for Web of Science ID 000320409100018

    View details for PubMedID 23254756

  • Food limited butterflies Resting and flight metabolic rate, fecundity and longevity Annual Meeting of the Society-for-Integrative-and-Comparative-Biology (SICB) Niitepold, K., Perez, A., Boggs, C. L. OXFORD UNIV PRESS INC. 2013: E156–E156
  • Comparative Effects of Adult Food Limitation on Butterfly Life Histories Annual Meeting of the Society-for-Integrative-and-Comparative-Biology (SICB) Boggs, C. L., Niitepold, K., Perez, A. OXFORD UNIV PRESS INC. 2013: E19–E19
  • A single climate driver has direct and indirect effects on insect population dynamics ECOLOGY LETTERS Boggs, C. L., Inouye, D. W. 2012; 15 (5): 502-508


    Weather drives population dynamics directly, through effects on vital rates, or indirectly, through effects on the population's competitors, predators or prey and thence on vital rates. Indirect effects may include non-additive interactions with density dependence. Detection of climate drivers is critical to predicting climate change effects, but identification of potential drivers may depend on knowing the underlying mechanisms. For the butterfly Speyeria mormonia, one climate driver, snow melt date, has multiple effects on population growth. Snow melt date in year t has density-dependent indirect effects. Through frost effects, early snow melt decreases floral resources, thence per-capita nectar availability, which determines fecundity in the lab. Snow melt date in year t + 1 has density-independent direct effects. These effects explain 84% of the variation in population growth rate. One climate parameter thus has multiple effects on the dynamics of a species with non-overlapping generations, with one effect not detectable without understanding the underlying mechanism.

    View details for DOI 10.1111/j.1461-0248.2012.01766.x

    View details for Web of Science ID 000302288900013

    View details for PubMedID 22414183

  • Native and Non-Native Community Assembly through Edaphic Manipulation: Implications for Habitat Creation and Restoration RESTORATION ECOLOGY Bonebrake, T. C., Navratil, R. T., Boggs, C. L., Fendorf, S., Field, C. B., Ehrlichl, P. R. 2011; 19 (6): 709-716
  • One variable species or multiple cryptic? Mitochondrial phylogeny of Central and North American Chlosyne lacinia (Lepidoptera: Nymphalidae) EUROPEAN JOURNAL OF ENTOMOLOGY Bonebrake, T. C., Watt, W. B., Perez, A., Boggs, C. L. 2011; 108 (4): 529-535
  • Population decline assessment, historical baselines, and conservation CONSERVATION LETTERS Bonebrake, T. C., Christensen, J., Boggs, C. L., Ehrlich, P. R. 2010; 3 (6): 371-378
  • More than just indicators: A review of tropical butterfly ecology and conservation BIOLOGICAL CONSERVATION Bonebrake, T. C., Ponisio, L. C., Boggs, C. L., Ehrlich, P. R. 2010; 143 (8): 1831-1841
  • Oviposition behavior and offspring performance in herbivorous insects: consequences of climatic and habitat heterogeneity OIKOS Bonebrake, T. C., Boggs, C. L., McNally, J. M., Ranganathan, J., Ehrlich, P. R. 2010; 119 (6): 927-934
  • Saliva or Regurgitated Nectar? What Heliconius Butterflies (Lepidoptera: Nymphalidae) Use for Pollen Feeding ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Eberhard, S. H., Hikl, A. L., Boggs, C. L., Krenn, H. N. 2009; 102 (6): 1105-1108
  • Does dietary restriction reduce life span in male fruit-feeding butterflies? EXPERIMENTAL GERONTOLOGY Molleman, F., Ding, J., Boggs, C. L., Carey, J. R., Arlet, M. E. 2009; 44 (9): 601-606


    Male life history and resource allocation is not frequently studied in aging and life span research. Here, we verify that males of long-lived fruit-feeding butterfly species have reduced longevity on restricted diets [Beck, J., 2007. The importance of amino acids in the adult diet of male tropical rainforest butterflies. Oecologia 151, 741-747], in contrast to the common finding of longevity extension in dietary restriction experiments in Drosophila and some other organisms. Males of some of the most long-lived species of fruit-feeding butterflies were collected from Kibale Forest, Uganda, and kept on diets of either sugar or mashed banana. Seven out of eight species had non-significantly longer life spans on mashed banana diets. Data analysis using a time-varying Cox-model with species as covariate showed that males had reduced survival on the sugar diet during the first 35 days of captive life, but the effect was absent or reversed at more advanced ages. These results challenge the generality of dietary restriction as a way to extend life span in animals. We argue that such studies on males are promising tools for better understanding life history evolution and aging because males display a wider variety of tactics for obtaining reproductive success than females.

    View details for DOI 10.1016/j.exger.2009.06.008

    View details for Web of Science ID 000270016700006

    View details for PubMedID 19580860

  • Understanding insect life histories and senescence through a resource allocation lens FUNCTIONAL ECOLOGY Boggs, C. L. 2009; 23 (1): 27-37
  • Longevity can buffer plant and animal populations against changing climatic variability ECOLOGY Morris, W. F., Pfister, C. A., Tuljapurkar, S., Haridas, C. V., Boggs, C. L., Boyce, M. S., Bruna, E. M., Church, D. R., Coulson, T., Doak, D. F., Forsyth, S., Gaillard, J., Horvitz, C. C., Kalisz, S., Kendall, B. E., Knight, T. M., Lee, C. T., Menges, E. S. 2008; 89 (1): 19-25


    Both means and year-to-year variances of climate variables such as temperature and precipitation are predicted to change. However, the potential impact of changing climatic variability on the fate of populations has been largely unexamined. We analyzed multiyear demographic data for 36 plant and animal species with a broad range of life histories and types of environment to ask how sensitive their long-term stochastic population growth rates are likely to be to changes in the means and standard deviations of vital rates (survival, reproduction, growth) in response to changing climate. We quantified responsiveness using elasticities of the long-term population growth rate predicted by stochastic projection matrix models. Short-lived species (insects and annual plants and algae) are predicted to be more strongly (and negatively) affected by increasing vital rate variability relative to longer-lived species (perennial plants, birds, ungulates). Taxonomic affiliation has little power to explain sensitivity to increasing variability once longevity has been taken into account. Our results highlight the potential vulnerability of short-lived species to an increasingly variable climate, but also suggest that problems associated with short-lived undesirable species (agricultural pests, disease vectors, invasive weedy plants) may be exacerbated in regions where climate variability decreases.

    View details for Web of Science ID 000253717200003

    View details for PubMedID 18376542

  • A trade-off between female lifespan and larval diet breadth at the interspecific level in Lepidoptera EVOLUTIONARY ECOLOGY Jervis, M. A., Ferns, P. N., Boggs, C. L. 2007; 21 (3): 307-323
  • When agendas collide: Human welfare and biological conservation CONSERVATION BIOLOGY Chan, K. M., Pringle, R. M., Ranganathan, J., Boggs, C. L., Chan, Y. L., Ehrlich, P. R., Haff, P. K., Heller, N. E., Al-Krafaji, K., Macmynowski, D. P. 2007; 21 (1): 59-68


    Conservation should benefit ecosystems, nonhuman organisms, and current and future human beings. Nevertheless, tension among these goals engenders potential ethical conflicts: conservationists' true motivations may differ from the justifications they offer for their activities, and conservation projects have the potential to disempower and oppress people. We reviewed the promise and deficiencies of integrating social, economic, and biological concerns into conservation, focusing on research in ecosystem services and efforts in community-based conservation. Despite much progress, neither paradigm provides a silver bullet for conservation's most pressing problems, and both require additional thought and modification to become maximally effective. We conclude that the following strategies are needed to make conservation more effective in our human-dominated world. (1) Conservation research needs to integrate with social scholarship in a more sophisticated manner. (2) Conservation must be informed by a detailed understanding of the spatial, temporal, and social distributions of costs and benefits of conservation efforts. Strategies should reflect this understanding, particularly by equitably distributing conservation's costs. (3) We must better acknowledge the social concerns that accompany biodiversity conservation; accordingly, sometimes we must argue for conservation for biodiversity's sake, not for its direct human benefits.

    View details for DOI 10.1111/j.1523-1739.2006.00570.x

    View details for Web of Science ID 000244148800014

    View details for PubMedID 17298511

  • Egg maturation strategy and survival trade-offs in holometabolous insects: a comparative approach BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY Jervis, M. A., Boggs, C. L., Ferns, P. N. 2007; 90 (2): 293-302
  • Egg maturation strategy and survival trade-offs in holometabolous insects: A comparative approach Biological Journal of the Linnean Society Jervis MA, Boggs CL, Ferns PN 2007; 90: 293-302
  • Delayed population explosion of an introduced butterfly JOURNAL OF ANIMAL ECOLOGY Boggs, C. L., Holdren, C. E., Kulahci, I. G., Bonebrake, T. C., Inouye, B. D., Fay, J. P., McMillan, A., Williams, E. H., Ehrlich, P. R. 2006; 75 (2): 466-475


    1. The causes of lagged population and geographical range expansions after species introductions are poorly understood, and there are relatively few detailed case studies. 2. We document the 29-year history of population dynamics and structure for a population of Euphydryas gillettii Barnes that was introduced to the Colorado Rocky Mountains, USA in 1977. 3. The population size remained low (< 200 individuals) and confined to a single habitat patch (approximately 2.25 ha) to 1998. These values are similar to those of many other populations within the natural geographical range of the species. 4. However, by 2002 the population increased dramatically to > 3000 individuals and covered approximately 70 ha, nearly all to the south of the original site. The direction of population expansion was the same as that of predominant winds. 5. By 2004, the butterfly's local distribution had retracted mainly to three habitat patches. It thus exhibited a 'surge/contraction' form of population growth. Searches within 15 km of the original site yielded no other new populations. 6. In 2005, butterfly numbers crashed, but all three habitat patches remained occupied. The populations within each patch did not decrease in the same proportions, suggesting independent dynamics that are characteristic of metapopulations. 7. We postulate that this behaviour results, in this species, in establishment of satellite populations and, given appropriate habitat structure, may result in lagged or punctuated expansions of introduced populations.

    View details for DOI 10.1111/j.1365-2656.2006.01067.x

    View details for Web of Science ID 000236384700015

    View details for PubMedID 16637999

  • Demography in an increasingly variable world Trends in Ecology & Evolution Boyce MS, Haridas CV, Lee C, Boggs CL, Bruna EM, Coulson T, Doak D, Drake JM, Gaillard J-M, Horvitz CC, Kalisz S, Kendall BE, Knight T, Menges ES, Morris WF, Pfister CA, Tuljapurkar SD 2006; 21: 141-147
  • Linking nectar amino acids to fitness in female butterflies TRENDS IN ECOLOGY & EVOLUTION Jervis, M. A., Boggs, C. L. 2005; 20 (11): 585-587


    Nectar of butterfly-pollinated flowers contains generally higher levels of amino acids than does nectar of flowers pollinated by most other animal types. One proposed explanation is that these amino acids promote butterfly fitness, although the evidence has been equivocal. In a new study, Mevi-Schütz and Erhardt showed that nectar amino acids enhanced fecundity in the butterfly Araschnia levana, but only when the larval diet was poor. Their results support the hypothesis that butterflies are agents of selection for higher nectar amino acid production, suggest that the larval food plant has a key role in the evolution of the flower-butterfly mutualism, and demonstrate that the importance, to butterfly reproduction, of different nutrient sources varies with butterfly nutritional state.

    View details for DOI 10.1016/j.tree.2005.08.015

    View details for Web of Science ID 000233213100004

    View details for PubMedID 16701440

  • Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY Stjernholm, F., Karlsson, B., Boggs, C. L. 2005; 86 (3): 363-380
  • The amino acids used in reproduction by butterflies: A comparative study of dietary sources using compound-specific stable isotope analysis PHYSIOLOGICAL AND BIOCHEMICAL ZOOLOGY O'Brien, D. M., Boggs, C. L., Fogel, M. L. 2005; 78 (5): 819-827


    It is a nutritional challenge for nectar-feeding insects to meet the amino acid requirements of oviposition. Here we investigate whether egg amino acids derive from larval diet or are synthesized from nectar sugar in four species of butterfly: Colias eurytheme, Speyeria mormonia, Euphydryas chalcedona, and Heliconius charitonia. These species exhibit a range of life history and differ in degree of shared phylogeny. We use 13C differences among plants to identify dietary sources of amino acid carbon, and we measure amino acid 13C using compound-specific stable isotope analysis. Egg essential amino acids derived solely from the larval diet, with no evidence for metabolic carbon remodeling. Carbon in nonessential amino acids from eggs derived primarily from nectar sugars, with consistent variation in amino acid turnover. There was no relationship between the nonessential amino acids of eggs and host plants, demonstrating extensive metabolic remodeling. Differences between species in carbon turnover were reflected at the molecular level, particularly by glutamate and aspartate. Essential amino acid 13C varied in a highly consistent pattern among larval host plants, reflecting a common isotopic "fingerprint" associated with plant biosynthesis. These data demonstrate conservative patterns of amino acid metabolism among Lepidoptera and the power of molecular stable isotope analyses for evaluating nutrient metabolism in situ.

    View details for Web of Science ID 000231502300014

    View details for PubMedID 16096984

  • Egg maturation strategy and its associated trade-offs: a synthesis focusing on Lepidoptera ECOLOGICAL ENTOMOLOGY JERVIS, M. A., Boggs, C. L., Ferns, P. N. 2005; 30 (4): 359-375
  • Larval food limitation in butterflies: effects on adult resource allocation and fitness OECOLOGIA Boggs, C. L., Freeman, K. D. 2005; 144 (3): 353-361


    Allocation of larval food resources affects adult morphology and fitness in holometabolous insects. Here we explore the effects on adult morphology and female fitness of larval semi-starvation in the butterfly Speyeria mormonia. Using a split-brood design, food intake was reduced by approximately half during the last half of the last larval instar. Body mass and forewing length of resulting adults were smaller than those of control animals. Feeding treatment significantly altered the allometric relationship between mass and wing length for females but not males, such that body mass increased more steeply with wing length in stressed insects as compared to control insects. This may result in changes in female flight performance and cost. With regard to adult life history traits, male feeding treatment or mating number had no effect on female fecundity or survival, in agreement with expectations for this species. Potential fecundity decreased with decreasing body mass and relative fat content, but there was no independent effect of larval feeding treatment. Realized fecundity decreased with decreasing adult survival, and was not affected by body mass or larval feeding treatment. Adult survival was lower in insects subjected to larval semi-starvation, with no effect of body mass. In contrast, previous laboratory studies on adult nectar restriction showed that adult survival was not affected by such stress, whereas fecundity was reduced in direct 11 proportion to the reduction of adult food. We thus see a direct impact of larval dietary restriction on survival, whereas fecundity is affected by adult dietary restriction, a pattern reminiscent of a survival/reproduction trade-off, but across a developmental boundary. The data, in combination with previous work, thus provide a picture of the intra-specific response of a suite of traits to ecological stress.

    View details for DOI 10.1007/s00442-005-0076-6

    View details for Web of Science ID 000231247800002

    View details for PubMedID 15891831

  • Flight muscle breakdown: A possible route to increased reproductive potential in butterflies? Biological Journal Linnean Society Stjernholm F, Karlsson B, Boggs CL 2005; 86: 363-380
  • Egg maturation strategy and its associated trade-offs: A synthesis focusing on Lepidoptera Ecological Entomology Jervis MA, Boggs CL, Ferns PN 2005; 30: 1-17
  • Allocation of larval and adult resources to reproduction in a fruit-feeding butterfly FUNCTIONAL ECOLOGY Fischer, K., O'Brien, D. M., Boggs, C. L. 2004; 18 (5): 656-663
  • Resource specialization in puddling Lepidoptera ENVIRONMENTAL ENTOMOLOGY Boggs, C. L., Dau, B. 2004; 33 (4): 1020-1024
  • Evolutionary genetics of dorsal wing colour in Colias butterflies JOURNAL OF EVOLUTIONARY BIOLOGY Ellers, J., Boggs, C. L. 2004; 17 (4): 752-758


    The evolution of butterfly wing colouration is strongly affected by its multiple functions and by the correlated evolution of wing colour elements. Both factors may prevent local adaptation to ecological conditions. We investigated one aspect of wing colouration, the degree of dorsal wing melanization, in the butterfly Colias philodice eriphyle across an elevational gradient and its correlation with another aspect of wing colouration, ventral wing melanization. Dorsal wing melanization increased with elevation and these differences persisted in a common environment. Full-sibling analysis revealed high heritability for males but only intermediate heritability for females. The correlation between ventral and dorsal melanization showed significant elevational and sex-specific differences. In males the two traits were highly correlated, whereas in females the strength of the correlation decreased with increasing elevation. We conclude that uncoupling of ventral and dorsal melanization has evolved in females but not in males and discuss possible mechanisms underlying uncoupling.

    View details for DOI 10.1111/j.1420-9101.2004.00736.x

    View details for Web of Science ID 000222052100004

    View details for PubMedID 15271074

  • Functional ecological implications of intraspecific differences in wing melanization in Colias butterflies BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY Ellers, J., Boggs, C. L. 2004; 82 (1): 79-87
  • Making eggs from nectar: the role of life history and dietary carbon turnover in butterfly reproductive resource allocation OIKOS O'Brien, D. M., Boggs, C. L., Fogel, M. L. 2004; 105 (2): 279-291
  • Functional ecological implications of intraspecific differences in wing melanization in Colias butterflies Biological Journal Linnean Society Ellers J, Boggs CL 2004; 82: 79-87
  • Making eggs from nectar: Connections between butterfly life history and the importance of nectar carbon in reproduction Oikos O'Brien DM, Boggs CL, Fogel ML 2004; 105: 279-291
  • Resource specialization in puddling Lepidoptera Environmental Entomology Boggs CL, Dau B 2004; 33: 1020-1024
  • Allocation of larval and adult resources to reproduction in a fruit-feeding butterfly Functional Ecology Fischer K, O'Brien DM, Boggs CL 2004; 18: 656-663
  • Pollen feeding in the butterfly Heliconius charitonia: isotopic evidence for essential amino acid transfer from pollen to eggs PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES O'Brien, D. M., Boggs, C. L., Fogel, M. L. 2003; 270 (1533): 2631-2636


    Heliconius and Laparus butterflies exhibit a unique pollen-collecting behaviour that enhances lifespan and fecundity. The specific nutritional contribution of pollen, however, had not been previously demonstrated. We used stable isotope variation to trace the carbon flow into eggs from corn pollen provided experimentally to ovipositing female Heliconius charitonia, and to evaluate the use of isotopically contrasting nectar sugars in egg amino acids. The delta(13)C of individual amino acids from pollen, larval host plant and the eggs from experimental butterflies was measured with gas chromatography/combustion/isotope ratio mass spectrometry (GC/C/IRMS), to evaluate amino acid transfer. The delta(13)C of egg essential amino acids indicated a transfer of essential amino acids from pollen to butterfly eggs. However, the delta(13)C of non-essential amino acids reflected the isotopic composition of the artificial nectar, indicating that H. charitonia synthesizes non-essential amino acids from dietary sugars. This, to our knowledge, is the first direct demonstration of amino acid transfer from pollen to butterfly eggs, and suggests that essential amino acids in particular are a key resource for extending lifespan and fecundity in Heliconius butterflies.

    View details for DOI 10.1098/rspb.2003.2552

    View details for Web of Science ID 000187989900014

    View details for PubMedID 14728787

  • The evolution of wing color: Male mate choice opposes adaptive wing color divergence in Colias butterflies EVOLUTION Ellers, J., Boggs, C. L. 2003; 57 (5): 1100-1106


    Correlated evolution of mate signals and mate preference may be constrained if selection pressures acting on mate preference differ from those acting on mate signals. In particular, opposing selection pressures may act on mate preference and signals when traits have sexual as well as nonsexual functions. In the butterfly Colias philodice eriphyle, divergent selection on wing color across an elevational gradient in response to the thermal environment has led to increasing wing melanization at higher elevations. Wing color is also a long-range signal used by males in mate searching. We conducted experiments to test whether sexual selection on wing melanization via male mate choice acts in the same direction as natural selection on mate signals due to the thermal environment. We performed controlled mate choice experiments in the field over an elevational range of 1500 meters using decoy butterflies with different melanization levels. Also, we obtained a more direct estimate of the relation between wing color and sexual selection by measuring mating success in wild-caught females. Both our experiments showed that wing melanization is an important determinant of female mating success in C. p. eriphyle. However, a lack of elevational variation in male mate preference prevents coevolution of mate signals and mate preference, as males at all elevations prefer less-melanized females. We suggest that this apparently maladaptive mate choice may be maintained by differences in detectability between the morphs or by preservation of species recognition.

    View details for Web of Science ID 000183500900014

    View details for PubMedID 12836826

  • Do hypotheses from short-term studies hold in the long-term? An empirical test ECOLOGICAL ENTOMOLOGY Hellmann, J. J., Weiss, S. B., McLaughlin, J. F., Boggs, C. L., Ehrlich, P. R., Launer, A. E., Murphy, D. D. 2003; 28 (1): 74-84
  • Countryside biogeography of tropical butterflies CONSERVATION BIOLOGY Horner-Devine, M. C., Daily, G. C., Ehrlich, P. R., Boggs, C. L. 2003; 17 (1): 168-177
  • Countryside biogeography of tropical butterflies Conservation Biology Horner-Devine MC, Daily GC, Ehrlich PR, Boggs CL 2003; 17: 168-177
  • Butterflies: Ecology and Evolution Taking Flight University of Chicago Press Boggs CL, Watt WB, Ehrlich PR 2003
  • Testing short-term hypotheses with a long-term study of a butterfly population Ecological Entomology Hellmann JJ, Weiss SB, McLaughlin JH, Boggs CL, Ehrlich PR, Launer AE, Murphy DD 2003; 28: 74-84
  • The route to extinction: population dynamics of a threatened butterfly OECOLOGIA McLaughlin, J. F., Hellmann, J. J., Boggs, C. L., Ehrlich, P. R. 2002; 132 (4): 538-548
  • Assessing the roles of patch quality, area, and isolation in predicting metapopulation dynamics CONSERVATION BIOLOGY Fleishman, E., Ray, C., Sjogren-Gulve, P., Boggs, C. L., Murphy, D. D. 2002; 16 (3): 706-716
  • Climate change hastens population extinctions PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA McLaughlin, J. F., Hellmann, J. J., Boggs, C. L., Ehrlich, P. R. 2002; 99 (9): 6070-6074


    Climate change is expected to alter the distribution and abundance of many species. Predictions of climate-induced population extinctions are supported by geographic range shifts that correspond to climatic warming, but few extinctions have been linked mechanistically to climate change. Here we show that extinctions of two populations of a checkerspot butterfly were hastened by increasing variability in precipitation, a phenomenon predicted by global climate models. We model checkerspot populations to show that changes in precipitation amplified population fluctuations, leading to rapid extinctions. As populations of checkerspots and other species become further isolated by habitat loss, climate change is likely to cause more extinctions, threatening both species diversity and critical ecosystem services.

    View details for DOI 10.1073/pnas.052131199

    View details for Web of Science ID 000175377800059

    View details for PubMedID 11972020

  • Renewable and nonrenewable resources: Amino acid turnover and allocation to reproduction in lepidoptera PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA O'Brien, D. M., Fogel, M. L., Boggs, C. L. 2002; 99 (7): 4413-4418


    The allocation of nutritional resources to reproduction in animals is a complex process of great evolutionary significance. We use compound-specific stable isotope analysis of carbon (GC/combustion/isotope ratio MS) to investigate the dietary sources of egg amino acids in a nectar-feeding hawkmoth. Previous work suggests that the nutrients used in egg manufacture fall into two classes: those that are increasingly synthesized from adult dietary sugar over a female's lifetime (renewable resources), and those that remain exclusively larval in origin (nonrenewable resources). We predict that nonessential and essential amino acids correspond to these nutrient classes and test this prediction by analyzing egg amino acids from females fed isotopically distinct diets as larvae and as adults. The results demonstrate that essential egg amino acids originate entirely from the larval diet. In contrast, nonessential egg amino acids were increasingly synthesized from adult dietary sugars, following a turnover pattern across a female's lifetime. This study demonstrates that female Lepidoptera can synthesize a large fraction of egg amino acids from nectar sugars, using endogenous sources of nitrogen. However, essential amino acids derive only from the larval diet, placing an upper limit on the use of adult dietary resources to enhance reproductive success.

    View details for Web of Science ID 000174856000052

    View details for PubMedID 11930002

  • The evolution of wing color in Colias butterflies: Heritability, sex linkage, and population divergence EVOLUTION Ellers, J., Boggs, C. L. 2002; 56 (4): 836-840


    We investigated the genetic background of intraspecific variation in wing color across an elevational gradient in the butterfly Colias philodice eriphyle. The degree of wing melanization was an accelerating function of elevation, and differences in wing melanization persisted in a common environment. Full-sibling analysis and parent-offspring regression yielded consistent, moderate to high heritabilities for the degree of wing melanization. The breeding experiments also demonstrated that wing melanization is strongly sex linked. Because traits that differentiate sister species also tend to be sex linked, our results suggest that the genetic mechanisms underlying intraspecific differences in wing melanization are not fundamentally different from those that have been shown to differentiate sister species.

    View details for Web of Science ID 000175614400018

    View details for PubMedID 12038541

  • The route to extinction: Population dynamics of a threatened butterfly Oecologia McLaughlin JF, Hellmann J, Boggs CL, Ehrlich PR 2002; 132: 538-548
  • Amino acid metabolism in adult Lepidoptera: from biochemistry to life history evolution. O'Brien, D. M., Fogel, M. L., Boggs, C. L. SOC INTEGRATIVE COMPARATIVE BIOLOGY. 2001: 1542–42
  • Physiological constraints on reproductive nutrient allocation in Lepidoptera: The dietary origins of individual egg amino acids. O'Brien, D. M., Boggs, C. L., Fogel, M. L. SOC INTEGRATIVE COMPARATIVE BIOLOGY. 1999: 95A–95A
  • Reproductive allocation from reserves and income in butterfly species with differing adult diets ECOLOGY Boggs, C. L. 1997; 78 (1): 181-191
  • Dynamics of reproductive allocation from juvenile and adult feeding: Radiotracer studies ECOLOGY Boggs, C. L. 1997; 78 (1): 192-202
  • Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies ECOLOGICAL ENTOMOLOGY Sculley, C. E., Boggs, C. L. 1996; 21 (2): 193-197
  • Rates of nectar feeding in butterflies: effects of sex, size, age and nectar concentration FUNCTIONAL ECOLOGY Boggs, C. L. 1988; 2 (3): 289-295


    Radiotracer studies on three butterfly species showed that nutrients contributed by males through mating are used by females for egg production and possibly for somatic maintenance.

    View details for Web of Science ID A1979HM63800036

    View details for PubMedID 17812454



    New oocytes are generated throughout long lives in butterflies of the genus Heliconius, which as adults feed on amino acids from pollen. In Dryas julia, a related heliconiine that feeds only on nectar and is relatively short-lived, the original oocyte supply is eventually depleted. Such divergent ovarian dynamics in closely related organisms are significant in terms of both their evolutionary basis and their physiological controls.

    View details for Web of Science ID A1977DN59900029

    View details for PubMedID 17783249